In Thailand and Five New Records for Thai Moss Flora Si HE1, Phiangphak

Hattoria 12: 27–39. 2021

Taxonomic updates and treatments of Hypnum (Hypnaceae) in Thailand and ﬁve new records for Thai moss ﬂora

Si HE1, Phiangphak SUKKHARAK2 & Sahut CHANTANAORRAPINT3

1 Missouri Botanical Garden, 4344 Shaw Blvd., St. Louis, MO 63110, U.S.A. 2 Department of Biology, Faculty of Science, Burapha University, Chonburi 20131, Thailand 3 Division of Biological Science, Faculty of Science, Prince of Songkla University, Hat Yai, Songkhla 90112, Thailand Author for correspondence: Si HE, [email protected]

Abstract A recent investigation into the four species of Hypnum Hedw. s. l. hitherto reported for Thailand confirms only two species that are consequently placed in Calohypnum Sakurai: Hypnum darjeelingense Ando [＝C. sikkimense (Renauld & Cardot) Jan Kučera & Ignatov] and H. plumiforme Wilson [＝C. plumiforme (Wilson) Jan Kučera & Ignatov]. The previous records of H. sakuraii (Sakurai) Ando and H. submolluscum Besch. in Thailand were based on misidentifications. The genus Hypnum is excluded from Thailand and Calohypnum is recognized. Based on the latest collections we made, five mosses: Claopodium aciculum (Broth.) Broth., Dicranoloma reflexum (Müll.Hal.) Renauld, Entodon scariosus Renauld & Cardot, Taxiphyllum giraldii (Müll.Hal.) M.Fleisch., and Vesicularia flaccida (Sull. & Lesq.) Z.Iwats. are reported new to Thai moss flora. The descriptions, illustrations, and key to species of Calohypnum in Thailand as well as diagnostic characters of the new moss records are provided.

Introduction Thailand has a diverse landscape, ranging from sea level to 2,550 meters in elevation. The most noticeable topographic features of the country include several high mountains, a lowland central plain dominated by the Chao Phraya River, and an undulating dry upland plateau bordered by the Mekong River. The main climate is characterized by tropical monsoon system which reverses its cycle seasonably. The distinctive climate and the variability in landscape have sustained a diverse plant vegetation that harbors a large number of bryophytes. With about 830 species, Thailand holds one of the richest moss ﬂoras in Southeast Asia (He 1998; Sukkharak & Chantanaorrapint 2014). Several recent bryological expeditions to Thailand supported by a National Geographic Society research grant uncovered numerous interesting mosses, including new records for the country. We have collected new specimens that might usually be identiﬁable to be species of Hypnum Hedw, namely H. darjeelingense Ando, also known as Ectropothecium sikkimense

27 (Renauld & Cardot) Renauld & Cardot, and H. plumiforme Wilson. Hypnum s. l. has a worldwide distribution tendency, but is much less common in the tropical and subtropical regions where Hypnum-like taxa were replaced by species of Ectropothecium Mitt. (He 2019). It is an uncommon genus in Thailand. The above-mentioned species were first reported from the country by an online checklist or in herbarium database without descriptions. Hypnum s. l. has traditionally been defined by a combination of the characters, i.e. prostrate and glossy plants, falcate-secund leaves with short double costae, well defined, subquadrate and thick-walled or enlarged oblong-rectangular, thin-walled alar cells, inclined to horizontal, cylindrical capsules, and perfect double peristomes. Until recently, the generic delimitation of Hypnum s. l. largely follows that of Ando (1972, 1973), which includes the type species of Calohypnum Sakurai, C. sakuraii Sakurai. Several phylogenetic studies indicate that Hypnum sensu Ando is clearly non-monophyletic and have since revised the structure of polyphyly of Hypnum in its traditional sense (Buck et al. 2000; Ignatov et al. 2007; Arikawa et al. 2008; Huttunen et al. 2012; Câmara et al. 2018; Schlesak et al. 2018). The new insight of polyphyly of Hypnum s. l. clearly supports a reclassification of the pertinent taxa that were traditional placed under the genus, resulting in recognition of additional genera outside the genus Hypnum. In accordance with well-supported molecular lineages as revealed by other similar studies on the phylogenetic revision of Hypnum, Kučera et al. (2019) proposed several taxonomic and nomenclatural re-arrangements, including resurrection of the generic status of Calohypnum. The molecular data suggest that numerous taxa in Ando’s subdivisions of Hypnum sect. Oldhamii Ando and sect. Curvifolia Ando form a clade separating from the Hypnum s. str. lineage that is typified by H. cupressiforme Hedw. A group of species formerly placed under Hypnum are now treated in Calohypnum, which is followed in this study. Nonetheless, no morphological traits can serve to separate Calohypnum from Hypnum. In this paper, we present a revised taxonomic account of the four Thai species that were previously named as Hypnum or under Ectropothecium and further recognize Calohypnum as a newly reported genus name for Thai moss flora. We also report Claopodium aciculum (Broth.) Broth., Dicranoloma reflexum (Müll.Hal.) Renauld, Entodon scariosus Renauld & Cardot, Taxiphyllum giraldii (Müll.Hal.) M.Fleisch., and Vesicularia flaccida (Sull. & Lesq.) Z.Iwats. as new moss records for the country based on our newly collected specimens.

Materials and Methods This study was based on the specimens collected by us and by C. Charoenphol and K. Larsen research team in Thailand. The voucher specimens were deposited at the Missouri Botanical Garden (MO), Prince of Songkla University (PSU), and in the Department of Biology, Faculty of Science, Burapha University (Hb. Burapha Univ.). A Nikon stereo zoom scope and a compound microscope (Nikon Eclipse E400) were used to examine specimens with standard anatomical and morphological methods applied to bryophytes. Microscopic images of morphological importance were captured with a digital camera (Nikon DS-U2) attached to the microscope.

28 Results & Discussion Calohypnum Sakurai, J. Jap. Bot. 25: 219. 1950. (Pylaisiaceae) Type: Calohypnum sakuraii Sakurai (lectotype designated by Kučera et al. 2019). Insomniella Hedenäs, Schlesak & D.Quandt, Bryophyte Diversity Evol. 40: 47. 2018.

Key to the species of Calohypnum in Thailand 1. Stem leaves ovate-lanceolate, distinctly falcate; leaf margins remotely serrulate at the upper half; costa of both stem and branch leaves thin, less than 1/5 the leaf length; median laminal cells ca. 3–4 µm wide, often smooth ············································· 1. C. plumiforme 1. Stem leaves triangularly lanceolate, slightly falcate; leaf margins strongly dentate or serrate at the upper half; costa of both stem and branch leave thick, up to 1/3 the leaf length; median laminal cells ca. 5–6 µm wide, usually prorate ························· 2. C. sikkimense

1. Calohypnum plumiforme (Wilson) Jan Kučera & Ignatov, Taxon 68: 651. 2019. Fig. 1. Basionym: Hypnum plumiforme Wilson, London J. Bot. 7: 277, pl. 10: d. 1848 (originally published as Hypnum plumaeforme). Type: China. From Chunshan to Hong Kong, Anderson s.n. (not seen). Insomniella plumiformis (Wilson) Hedenäs, Schlesak & D.Quandt, Bryoph. Diversity & Evol. 40: 47. 2018.

Plants moderately large, yellowish green or greenish, sometimes brownish. Stems creeping, up to 10 cm long, usually regularly pinnately branched; branches flattened or subjulaceous, ca. 1.5 cm long; in stem cross section central strand slightly developed; pseudoparaphyllia lanceolate mixed with narrowly foliose ones. Stem leaves broadly elliptic- to subcordate-lanceolate, branch leaves oblong-lanceolate, (1.4–)1.8–2.5(–3.0)×0.5–0.8(–1.0) mm; both leaves gradually acuminate, falcate-secund, plicate above; margins plane, serrulate above; costae double, short; upper and median leaf cells linear, 40–60(–70)×3–4 µm, somewhat thick-walled, sometimes prorate; basal cells shorter, thick-walled, more or less porose, yellowish brown; alar cells inflated, numerous, thin-walled, hyaline or yellowish, with 2–4 rows of smaller, subquadrate cells above. Dioicous. Perichaetial leaves erect, broadly ovate-lanceolate, long-acuminate, plicate, margins plane, serrulate, costae indistinct. Sporophyte not seen. Habitat: Forming a creeping patch on rocks and rocky soil in forest. Distribution: China, Japan, Korea, Russian Far East, India, Nepal, Thailand, Vietnam, the Philippines, Georgia (Caucasus, probably introduced, Ignatov et al. 2006), and also Hawaii Islands (Crum & Mueller-Dombois 1968, Ando 1972). Specimens examined: THAILAND. Chanthaburi: Khao Khitchakut National Park, 12°50′29″N, 102°09′45″E, 850–1,070 m, He & Sukkharak 47043, 47051 (both in Hb. Burapha Univ., MO). Songkhla: Ton Nga Chang Wildlife Sanctuary, 06°56′52″N, 100°14′01″E, 140–370 m elev., He & Sukkharak 47413 (Hb. Burapha Univ., MO). Taxonomic notes: The species was first reported from Thailand by Printarakul et al. (2009) in a database of bryophytes from the Chiang Mai University herbarium (CMUB), but no specimen was cited. This species often recognized as Hypnum species appears to be

29 Figure 1. Calohypnum plumiforme (Wilson) Jan Kučera & Ignatov. A, B. Stem leaves. C, D. Branch leaves. E. Habit. F. Pseudoparaphyllia, G. Leaf apex. H, I. Basal and alar leaf cells. J. Median leaf cells. Illustrations prepared from He & Sukkharak 47413 (MO).

somewhat widely distributed in East and Southeast Asia.

2. Calohypnum sikkimense (Renauld & Cardot) Jan Kučera & Ignatov, Taxon 68: 1383. 2020 (“2019”; 12 Mar. 2020). Fig. 2. Basionym: Hypnum hamulosum Schimp. var. sikkimense Renauld & Cardot, Bull. Soc.

30 Figure 2. Calohypnum sikkimense (Renauld & Cardot) Jan Kučera & Ignatov. A, B. Stem leaves. C, D. Branch leaves. E. Habit. F. Pseudoparaphyllia. G. Leaf apex. H. Basal leaf cells. I. Alar cells. J. Median leaf cells. Illustrations prepared from Sukkharak et al. 314 (MO).

Roy. Bot. Belgique 34: 78. 1895. Type: India. Sikkim, Darjeeling, Stevens s.n. (not seen). Calohypnum darjeelingense (Ando) Jan Kučera & Ignatov, Taxon 68: 651. 2019, illegitimate, type of earlier name included. Ectropothecium sikkimense (Renauld & Cardot) Renauld & Cardot, Bull. Soc. Roy. Bot. Belgique 41: 109. 1905.

31 Hypnum darjeelingense Ando, Hikobia 6: 41. 1971.

Plants fairly robust, yellowish green, glossy in low tufts. Main stems creeping, regularly pinnately branched in one plane (frondose), branch tips often hamate; pseudoparaphyllia numerous, lanceolate to narrowly foliose. Stem and branch leaves erectopatent, slightly falcate to second at tips, appressed when dry, triangularly lanceolate, narrowed at base, acuminate at apex, 1.1–1.2×0.30–0.35 mm; margin distinctly dentate at tips; costae double, thick, up to 1/3 the leaf length; laminal cells linear-rhomboidal, 50–60×5–6 µm, usually projecting at cell tips; alar cells distinctly differentiated, consisting of a small group of inflated cells and several rows of short-quadrate cells. Dioicous. Perichaetial leaves erect-spreading, triangularly lanceolate, long-acuminate, not plicate, margins serrulate, costae absent. Setae slender, ca. 1.5 cm long, often arcuate at apex. Capsules horizontal, cylindrical, cernuous, ca. 1.2×0.5 mm. Opercula relatively large, conic, shortly rostrate, ca. 0.7 mm long. Annuli unknown; Peristome hypnoid. Spores unknown. (Sporophytic features based on non-Thai collections). Habitat: On rocks. Distribution. India (Gangulee 1980), Nepal (Higuchi & Takaki 1988) and Thailand. Specimens examined: THAILAND. Chanthaburi: Khao Soi Dao, 900–1675 m elev., Sukkharak et al. 215, 314 (both in Hb. Burapha Univ., MO). Nakhon Ratchasima: Pak Thong Chai, Larsen et al. 3211 (AAU, MO); Khao Yai National Park, 14°29′N, 101°30′E, 1,000– 1,100 m elev., Charoenphol et al. 4311 (AAU, MO). Phang-nga: Nam Tok Pee, 08°15′N, 98°40′E, 50 m elev., Larsen et al. 31189 (AAU, MO). Ranong: Khao Phra Mi, 09°17′N, 98°26′E, 60 m elev., Larsen et al. 30808 (AAU, MO). Satun: Nam Tok Baripath, 07°02′N, 100°11′E, Charoenphol et al. 3914 (AAU, MO). Trang: Chao Pa Waterfall, 07°14′24″N, 99°50′38″E, 20–120 m elev., He & Sukkharak 47444 (Hb. Burapha Univ., MO). Trat: Ko Kut, 12°35′N, 101°31′E, Charoenphol et al. 5124 (AAU, MO). Taxonomic notes: The concept of Hypnum s. l. as circumscribed by Ando (1972, 1973) is rather broad. According to Ignatov et al. (2007), Huttunen et al. (2012), and Kučera et al. (2019), only those members of Hypnum sect. Hypnum in the sense of Ando were retained in their delimitation of Hypnum s. str. which is typified by H. cupressiforme Hedw. The genus Calohypnum now consists of six species that are predominantly distributed in East-Asia. Molecularly, the genus shares close affinity with primarily paleotropical Ectropothecium and Vesicularia (Müll.Hal.) Müll.Hal. (Kučera et al. 2019).

Hypnum species excluded from Thailand Hypnum sakuraii (Sakurai) Ando and H. submollucum Besch. were reported in an online moss checklist of Thailand (He 1998, http://www.mobot.org/MOBOT/moss/Thailand/) based on a collection from Khao Yai-Natioal Park, Prachinburi province (Larsen et al. 89, MO, as H. sakuraii) and a collection from Chatvarin Fall, Narathiwat province (Charoenphol et al. 4014, MO, as H. submollucum). The present study reveals that both specimens were misidentified. The collection (Larsen et al. 89) is a mixed specimen containing a part of Calohypnum plumiforme and a part of Ectropothecium sp. The collection (Charoenphol et al. 4014) also belongs to the genus Ectropothecium. The genus Ectropothecium is characterized by having a single inflated alar cell at a hardly differentiated alar region while Calohypnum usually has a

32 clearly diﬀerentiated alar region that often consists of a few enlarged alar cells.

New records to Thailand Based on collections made by us in our recent field trips, the following five mosses are herein reported new to Thailand. 1. Claopodium aciculum (Broth.) Broth., Nat. Pflanzenfam. I (3): 1009. 1908. Fig. 3. Basionym: Thuidium aciculum Broth., Hedwigia 38: 245. 1899.

This species is characterized by having more or less similar stem and branch leaves with percurrent costa and centrally unipapillose leaf cells. It is very close to rather widespread Claopodium prionophyllum (Müll.Hal.) Broth. in Thailand, but diﬀers by having smaller plants and elongate-oval to rhomboidal leaf cells and by its restricted distribution in East Asia. Habitat: On tree trunks. Distribution: China, Korea, Japan, Laos, and new to Thailand. Specimen examined: THAILAND. Loei: Phu Luang Wildlife Sanctuary, 17°20′02″N, 101°30′33″E, 915 m elev., He & Sukkharak 46904 (Hb. Burapha Univ., MO).

Figure 3. Claopodium aciculum (Broth.) Broth. A, B. Habits. C, D. Secondary stem leaves. E. Leaf apex. F, G. Upper leaf cells. H. Median leaf cells. I. Basal leaf cells. Illustrations prepared from He & Sukkharak 46904 (MO).

33 Figure 4. Dicranoloma reﬂexum (Müll.Hal.) Renauld. A, B. Habits. C, D. lower half leaves. E, F. Leaf apices. G. Upper leaf cells. H. Basal leaf cells. I. Alar cells. Illustrations prepared from He & Chantanaorrapint 47794 (MO).

2. Dicranoloma reﬂexum (Müll.Hal.) Renauld, Essai Leucoloma 14. 1909. Fig. 4. Basionym: Dicranum reﬂexum Müll.Hal., Syn. Musc. Frond. 1: 373. 1848.

The species is a rather large and beautiful moss (Klazenga 1999). It is characterized by the plicate leaves with two long, deep folds parallel to either side of the costa. It diﬀers from D. blumei (Nees) Broth. ex Renauld by having more distinct abaxial teeth on low ribs (vs scattered) and much wider costa (40–120 vs 20–50 µm) with more guide cells (4–11 vs 2–4 cells). The collections from Thailand were conﬁrmed by Dr. Klazenga when he visited the Missouri Botanical Garden in 2018. Habitat: On humus, base of trees, or on rocks. Distribution: Indonesia, Malaysia, the Philippines, and new to Thailand. Specimens examined: THAILAND. Loei: Phu Luang Wildlife Sanctuary, 17°16′13″N, 101°31′20″E, 1480–1510 m elev., He & Sukkharak 46741 (Hb. Burapha Univ., MO), He & Chantanaorrapint 47794, 47828a (both at MO, PSU).

3. Entodon scariosus Renauld & Cardot, Bull. Soc. Roy. Bot. Belgique 34: 75. 1896. Fig. 5. Cylindrothecium scariosum (Renauld & Cardot) Paris, Index Bryol. Suppl. 110. 1900.

34 Figure 5. Entodon scariosus Renauld & Cardot. A. Habit. B. Perchaetium. C. Perigonium. D. Perichaetial leaf. E, F. Leaves. G. Apical leaf cells. H. Basal leaf cells. Illustrations prepared from He & Sukkharak 47127 (MO).

Plants of this species are usually robust, up to 10 cm long. It diﬀers from other Entodon species from Thailand, such as E. macropodus (Hedw.) Müll.Hal. and E. schensianus Müll. Hal. by the narrowly and gradually acute leaf apices. When sporophytes are present, the species is characterized by the absence of annulus on its capsules and the yellowish setae that are unlike to usual reddish color of many Entodon species. Habitat: On rock. Distribution: China, India, and new to Thailand. Specimen examined: THAILAND. Prachuap Khiri Khan: Kaeng Krachan National Park, 12°47′54″N, 99°27′22″E, 270–290 m elev., He & Sukkharak 47127 (Hb. Burapha Univ., MO).

4. Taxiphyllum giraldii (Müll.Hal.) M.Fleisch., Musci Buitenzorg 4: 1435. 1923. Fig. 6. Basionym: Plagiothecium giraldii Müll. Hal., Nuovo Giorn, Bot. Ital., n.s. 3: 114. 1896. Isopterygium giraldii (Müll. Hal.) Paris, Index Bryol. Suppl. 219. 1900.

This species is distinguished by the closely complanate leafy stems. It is close to Taxiphyllum taxirameum (Mitt.) M.Fleisch., but diﬀers in having stem and branch leaves that are erect-spreading and in having shortly acute to obtuse branch leaves with distinctly dentate upper margins. In contrast, T. taxirameum has stem and branch leaves that are obliquely spreading and has gradually and narrowly acute branch leaves with ﬁnely serrulate upper margins. Habitat: On tree trunks. Distribution: China, Japan, and new to Thailand.

35 Figure 6. Taxiphyllum giraldii (Müll.Hal.) M.Fleisch. A. Habit. B. Pseudoparaphyllia. C, D. Stem leaves. E, F. Branch leaves. G. Apical leaf cells. H. Basal leaf cells. Illustrations prepared from He & Sukkharak 47372 (MO).

Specimen examined: THAILAND. Songkhla: Khao Nam Khang National Park, Ton Lad waterfall, 06°35′56″N, 100°35′18″E, 95–130 m elev., He & Sukkharak 47372 (Hb. Burapha Univ., MO).

5. Vesicularia flaccida (Sull. & Lesq.) Z.Iwats., J. Hattori Bot. Lab. 26: 70. 1963. Fig. 7. Basionym: Hypnum flaccidum Sull. & Lesq., Proc. Amer. Acad. Arts 4: 280. 1859. Isopterygium flaccidum (Sull. & Lesq.) Mitt., Trans. Linn. Soc. London, Bot. 3: 176. 1891.

Vesicularia ﬂaccida was originally described from Japan. It is uncommon throughout its distribution range. The species is diﬀerent from other species of Vesicularia from Thailand by having irregularly and sparsely branched stems and ovate-lanceolate leaves with abruptly tapering acumen and entire leaf margins. Habitat: On soil. Distribution: China, Japan, Korea, Russian Far East, and new to Thailand. Specimen examined: THAILAND. Prachuap Khiri Khan: Huai Yang Waterfall National Park, 11°38′52″N, 99°34′56″E, 1078–1204 m elev., He & Sukkharak 47238 (Hb. Burapha Univ., MO).

36 Figure 7. Vesicularia ﬂaccida (Sull. & Lesq.) Z.Iwats. A. Habit (capsule should be horizontal, here due to seta twisting just below the capsule). B, C. Leaves. D. Capsule. E. Apical leaf cells. F. Median leaf cells. G. Basal leaf cells. H. Exothecial cells of capsule. Illustrations prepared from He & Sukkharak 47238 (MO).

Acknowledgments We thank B. Hassama, C. Kraithep, W. Juengprayoon, P. Rojchana-umpawan, N. Putthisawong, J. Sangrattanaprasert, and O. Suwanmala for their helps during the ﬁeld work. We are grateful to C. Charoenphol, K. Larsen, I. Nielsen, T. Santisuk, T. Smitinand, A. Touw, and E. Warncke who collected specimens of mosses from Thailand and provided them to the Missouri Botanical Garden herbarium (MO), making this study possible. We are grateful to Dr. M. Ignatov and another anonymous reviewer for their helpful comments and important information about the species distribution. This research was supported by the National Geographic Society grant (NGS 9777-15).

References Ando H. 1972. Studies on the genus Hypnum (I). Journal of Science of the Hiroshima University, Series B, Division 2 (Botany) 14: 53–73. Ando H. 1973. Studies on the genus Hypnum Hedw. (II). Journal of Science of the Hiroshima University,

37 Series B, Division 2 (Botany) 14: 165–207. Arikawa T., Tsubota H., Deguchi H., Nishimura N. & Higuchi M. 2008. Phylogenetic analysis of the family Hypnaceae based on rbcL gene sequences. Pages 215–225. In: Mohamed H., Bakar B. B., Boyce A. N. & Yuen P. L. K. (eds.), Bryology in the New Millennium. Kuala Lumpur: Institute of Biological Sciences, University of Malaya. Buck W. R., Goffinet B. & Shaw A. J. 2000. Testing molecular concepts of orders of pleurocarpous mosses (Bryophyta) using phylogenetic reconstructions based on trnL-trnF and rps4 sequences. Molecular Phylogenetics and Evolution 16: 180–198. Câmara P. E. A. S., Carvalho-Silva M., Henriques D. K., Guerra J., Gallego M. T., Poveda D. R. & Stech M. 2018. Pylaisiaceae Schimp. (Bryophyta) revisited. Journal of Bryology 40: 251–264. Crum H. A. & Mueller-Dombois D. 1968. Two new mosses from Hawaii. The Journal of the Hattori Botanical Laboratory 31: 293–296. Gangulee H. C. 1980. Mosses of Eastern India and Adjacent Regions 8 (Hypnobryales–Hypnineae: Floristic trends: Index: Errata): pp. (xlv–liii)＋1753–2145. Privately published, Calcutta. He S.1998. An annotated checklist and atlas of the mosses of Thailand. http://www.mobot.org/MOBOT/ moss/Thailand/. Missouri Botanical Garden, St. Louis. Accessed: on April 30, 2021. He S. 2019. Disentangling the taxonomy and morphology of Ectropothecium (Bryophyta: Hypnaceae s. l.) with comments on its generic relationships, and Neoptychophyllum, a new name for the genus Ptychophyllum. Bryophyte Diversity and Evolution 41: 17–34. Higuchi M. & Takaki N. 1988. A collection of the hypnaceous mosses from the Kathmandu area, central Nepal. Pages 171–175. In: Watanabe M. & Malla S. B. (eds.), Cryptogams of the Himalayas. Vol. 1. The Kathmandu Valley. National Science Museum, Tsukuba. Huttunen S., Bell N., Bobrova V. K., Buchbender V., Buck W. R., Cox C. J., Goffinet B., Hedenäs L., Ho B.-C., Ignatov M. S., Krug M., Kuznetsova O. I., Milyutina I. A., Newton A. E., Olsson S., Pokorny-Montero L., Shaw J. A., Stech M., Troitsky A. V., Vanderpoorten A. & Quandt D. 2012. Disentangling knots of rapid evolution: Origin and diversification of the moss order Hypnales. Journal of Bryology 34: 187–211. Ignatov M. S., Afonina O. M. & Ignatova E. A. 2006. Check-list of mosses of East Europe and North Asia. Arctoa 15: 1–130. Ignatov M. S., Gardiner A. A., Bobrova V. K., Milyutina I. A., Huttunen S. & Troitsky A. V. 2007. On the relationships of mosses of the order Hypnales, with special reference to taxa traditionally classified in the Leskeaceae. In: Newton A. E. & Tangney R. S. (eds.), Pleurocarpous mosses: systematics and evolution. Systematics Association Special Volume 71: 177–213. Klazenga N. 1999. A revision of the Malesian species of Dicranoloma (Dicranaceae, Musci). The Journal of the Hattori Botanical Laboratory 87: 1–130. Kučera J., Kuznetsova O. I., Manukjanová A. & Ignatov M. S. 2019. A phylogenetic revision of the genus Hypnum: towards completion. Taxon 68: 628–660. Printarakul N., Wongkuna K., Kornochalert S. & Santanachote K. 2009. Database of bryophytes and their ecological parameters in the CMU Herbarium. Maejo International Journal of Science and Technology 3: 99–129. Schlesak S., Hedenäs L., Nebel M. & Quandt D. 2018. Cleaning a taxonomic dustbin: Placing the European Hypnum species in a phylogenetic context! Bryophyte Diversity Evolution 40: 37–54. Sukkharak P. & Chantanaorrapint S. 2014. Bryophyte studies in Thailand: past, present, and future.

38 Cryptogamie, Bryologie 35: 5–17. manuscript received April 19, 2021; accepted May 18, 2021